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(Decapoda, Brachyura, Mictyridae) from the Ryukyu Islands, Japan

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(Decapoda, Brachyura, Mictyridae) from the Ryukyu Islands, Japan A NEW SPECIES OF MICTYRIS (DECAPODA, BRACHYURA, MICTYRIDAE) FROM THE RYUKYU ISLANDS, JAPAN BY PETER J. F. DAVIE1,4), HSI-TE SHIH2,5) and BENNY K. K. CHAN3,6) 1) Queensland Museum, P.O. Box 3300, South Brisbane, Queensland, 4101, Australia 2) Department of Life Science, National Chung Hsing University, 250 Kuo-Kuang Road, Taichung 402, Taiwan 3) Research Centre for Biodiversity, Academia Sinica 128 section 2, Academia Road, Taipei 115, Taiwan ABSTRACT The Asian soldier crab Mictyris brevidactylus Stimpson, 1858, was re-examined genetically and morphologically from specimens collected across its range. It was found to consist of two cryptic sister species, with crabs from the Ryukyu Islands representing a new species, M. guinotae. Mictyris guinotae is smaller at maturity than M. brevidactylus, has plain blue colouring without red banding on the legs, and has a differently shaped apex of its male first gonopod. The two species also differ genetically, with 16S rRNA and COI showing a mean divergence (K2P distance) of 1.08% and 4.39%, respectively, representing 2.0 and 3.5 times more divergence than present intraspecifically. Results provide further evidence that some elements of the marine fauna of the Ryukyu Islands have had a separate evolutionary history from continental Asia as well as Taiwan. RÉSUMÉ Le crabe soldat asiatique Mictyris brevidactylus Stimpson, 1858, a été réexaminé à la fois génétiquement et morphologiquement à travers son aire de répartition. Il s’avère qu’il comprend deux espèces sœurs dont les crabes des îles Ryükyü représentent une nouvelle espèce, M. guinotae. M. guinotae est plus petit à la maturité que M. brevidactylus, il a une couleur bleue uniforme sans bandes rouges sur les pattes et la forme de l’extrémité du premier gonopode mâle est différente. Les deux espèces diffèrent aussi génétiquement avec le 16S rRNA et le COI montrant une divergence moyenne respective (distance K2P) de 1,08% et de 4,39%, ce qui représente 2 et 3,5 fois plus que la divergence intra spécifique. Les résultats fournissent de 4) e-mail: [email protected] 5) e-mail: [email protected] 6) Corresponding author; e-mail: [email protected] © Koninklijke Brill NV, Leiden, 2010 Studies on Brachyura: 83-105 84 CRM 011 – Castro et al. (eds.), BRACHYURA: A HOMAGE TO DANIÈLE GUINOT nouvelles preuves que quelques éléments de la faune marine des îles Ryükyü ont eu une histoire évolutive séparée de celle du continent asiatique, Taïwan compris. INTRODUCTION Soldier crabs (species of Mictyris Latreille, 1806) are a conspicuous inter- tidal crab on many shores throughout tropical and temperate Australia, South- east Asia and East Asia. They play an important ecological role in maintaining the proper functioning of the intertidal flats where they occur (Quinn, 1986; Dittman, 1994; Webb & Eyre, 2004) and experimental removal of one species, M. longicarpus Latreille, 1806, has resulted in the overgrowth of dense micro- bial mats (Webb & Eyre, 2004). Mictyris armies are also an important food source for shore birds (Zharikov & Skilleter, 2002, 2003, 2004; Rohweder & Lewis, 2004; Webb & Eyre, 2004). The family Mictyridae contains a single genus Mictyris,with,atpresent, five recognized species: M. longicarpus; M. platycheles H. Milne Edwards, 1852; M. brevidactylus Stimpson, 1858; M. livingstonei McNeill, 1926; and M. occidentalis Unno, 2008. Mictyris species are remarkably similar in their appearance, and this has caused confusion about their specific status. Despite McNeill’s (1926) excellent revision of the genus, there has been some longstanding confusion about the status of M. longicarpus. Mictyris longicarpus has been considered the only widespread species, with many literature records from throughout the tropical central Indo-West Pacific region (Takeda, 1978: 31). The first author is currently undertaking a modern revision of the genus, and after examining samples of Mictyris from throughout its known range, has concluded that M. longicarpus sensu stricto is confined to the eastern coast of Australia from where it was originally described. A number of new cryptic species, all with restricted, relatively narrow, geographical distributions will be described (P. J. F. Davie, in progress). Davie (2002) also pointed out that there were two undescribed species in northern and western Australia, with one of these, M. occidentalis, having been recently described by Unno (2008). Takeda (1978) firmly established the separate species status of Mictyris brevidactylus, and this paper should be referred to for a thorough analysis of the differences between that species and M. longicarpus.However,asweshall demonstrate in the present paper, the Ryukyu Islands material that Takeda assumed was characteristic of Mictyris brevidactylus elsewhere in Taiwan, China, and the Philippines, is in fact a separate species, which is here described as new. Davie et al., MICTYRIS GUINOTAE NOV. 85 MATERIAL AND METHODS The male first gonopods are referred to as G1. Measurements are given in millimetres. Carapace length (CL) was measured in the mid-line from the mid- dle of the rostrum to the posterior margin; carapace width (CW) was measured across the widest point of the carapace. The material examined is deposited in the Queensland Museum (QM), the Muséum national d’Histoire naturelle, Paris (MNHN), the Zoological Reference Collection (ZRC) of the Raffles Mu- seum of Biodiversity Research, National University of Singapore; Ryukyus University Museum, Okinawa (RUMF); National Museum of Natural Sci- ence (NMNS), Taichung, Taiwan; Zoological Collections of the Department of Life Science (NCHUZOOL), National Chung Hsing University, Taichung, Tai- wan; Coastal Ecology Laboratory, Academia Sinica, Taipei (CEL); Sencken- berg Museum, Frankfurt am Main (SMF); Western Australian Museum, Perth (WAM). Specimens of Mictyris for genetic analyses were collected from the Ryukyus, Taiwan, Hong Kong, and Hainan Island, China. Samples were pre- served in 75-95% ethanol after collection, and deposited in several museums (as above) (table I). Genomic DNA was isolated from the muscle tissue of legs using the DNA genomic extraction kit. A region of ∼550 base pairs (bp) at the 5-end of the mitochondrial 16S rRNA gene was selected for amplifica- tion by a polymerase chain reaction (PCR) using the primers, 1471 (5- CCTGTTTANCAAAAACAT-3) and 1472 (5-AGATAGAAACCAACCT GG-3) (Crandall & Fitzpatrick, 1996). A portion of the COI gene was ampli- fied by PCR using the primers, LCO1490 (5-GGTCAACAAATCATAAAGAT ATTGG-3) and HCO2198 (5-TAAACTTCAGGGTGACCAAAAAATCA- 3) (Folmer et al., 1994). PCR conditions for the above primers included de- naturation for 50 s at 94◦C, annealing for 70 s at 45◦C, and extension for 60 s at 72◦C, followed by a final extension for 10 min. at 72◦C. Sequences were ob- tained by automated sequencing (ABI PRISM 377 Sequencer and MegaBACE DNA Analysis System 500, Amersham, U.K.) and were aligned with the aid of CLUSTAL W (vers. 1.4, Thompson et al., 1994) and BIOEDIT (vers. 5.09, Hall, 2001), after verification with the complementary strand. Sequences of the different haplotypes were deposited in the DNA Data Bank of Japan (DDBJ), and the accession numbers are given in table I. The Kimura (1980) 2-parameter distance (K2P distance) between haplotypes was calculated by the software MEGA 4 (Tamura et al., 2007). This distance measure is adopted because it has been widely used for genetic comparisons among taxa (see Lefébure et al., 2006; Costa et al., 2007; Hubert et al., 2008). 86 CRM 011 – Castro et al. (eds.), BRACHYURA: A HOMAGE TO DANIÈLE GUINOT TABLE I Seven haplotypes of 16S ribosomal (r)RNA and 12 haplotypes of cytochrome oxidase subunit I (COI) genes of the two species of Mictyris from East Asia used in this study. NCHUZOOL, the Zoological Collections of the Department of Life Science, National Chung Hsing Univ.; CEL, Coastal Ecology Laboratory, Academia Sinica; DDBJ, the DNA Data Bank of Japan Species Localities Catalog no. Haplotypes DDBJ Haplotypes DDBJ of 16S access. no. of COI access. no. M. brevidactylus Waziwei, Taipei County, Taiwan CEL-MIC-TW2 Mb1 AB513630 Mb-C1 AB513638 Stimpson, 1858 Waziwei, Taipei County, Taiwan CEL-MIC-TW1 Mb2 AB513631 Mb-C2 AB513637 Siangshan, Hsinchu City, Taiwan CEL-MIC-TW3 Mb1 AB513630 Mb-C1 AB513638 Siangshan, Hsinchu City, Taiwan CEL-MIC-TW4 Mb1 AB513630 Mb-C1 AB513638 Haomeiliao, Chiayi County, Taiwan NCHUZOOL 13262 Mb1 AB513630 Mb-C1 AB513638 Starfish Bay, Hong Kong CEL-MIC-HK-1 Mb2 AB513631 Mb-C2b AB513639 Starfish Bay, Hong Kong CEL-MIC-HK-2 Mb2 AB513631 Mb-C2c AB513640 Wenchang, Hainan, China CEL-MIC-HN-1 Mb2 AB513631 Mb-C2d AB513641 Wenchang, Hainan, China CEL-MIC-HN-2 Mb2 AB513631 Mb-C2e AB513642 Wenchang, Hainan, China CEL-MIC-HN-3 Mb2 AB513631 Mb-C2f AB513643 Sanya, Hainan, China NCHUZOOL 13263 Mb2 AB513631 Mb-C2f AB513643 M. guinotae sp. nov. Awase, Okinawa, Ryukyus, Japan NCHUZOOL 13264 Mg1 AB513632 Mg-C1 AB513644 southern Naha, Okinawa, Ryukyus, Japan CEL-MIC-OK-1 Mg2 AB513633 Mg-C2 AB513645 southern Naha, Okinawa, Ryukyus, Japan CEL-MIC-OK-3 Mg2 AB513633 Mg-C2b AB513639 southern Naha, Okinawa, Ryukyus, Japan CEL-MIC-OK-2 Mg3 AB513634 Mg-C3 AB513646 Funaura Bay, Iriomote, Ryukyus, Japan NCHUZOOL 13266 Mg4 AB513635 Mg-C4 AB513647 Funaura Bay, Iriomote, Ryukyus, Japan NCHUZOOL 13266 Mg5 AB513636 Mg-C2b AB513648 Davie et al., MICTYRIS GUINOTAE NOV. 87 GENETIC RESULTS A 553 bp segment (excluding the primer regions) of the 16S rRNA from all 17 specimens was amplified and aligned; 12 positions were variable and 5 parsimoniously informative. Among the total number of sequences, 7 different haplotypes were distinguished (table I). The studied segment of 16S rRNA sequences was AT rich (71.8%) (T, 34.5%; A, 37.3%; G, 17.7%; and C, 10.5%). For the COI gene, a 658 bp segment was compared, resulting in 12 different haplotypes (table I). The studied segment of the COI sequence was also AT rich (63.0%) (T, 39.1%; A, 23.9%; G, 17.8%; and C, 19.2%).
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